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14 Fiorelli.Pmd Arquivos do Museu Nacional, Rio de Janeiro, v.65, n.4, p.417-459, out./dez.2007 ISSN 0365-4508 THE FIRST “PROTOSUCHIAN” (ARCHOSAURIA: CROCODYLIFORMES) FROM THE CRETACEOUS (SANTONIAN) OF GONDWANA 1 (With 16 figures) LUCAS E. FIORELLI 2 JORGE O. CALVO 3 ABSTRACT: The remains of “protosuchians” from the Cretaceous come, to exception of “Las Hoyas crocodyliform” from the Lower Cretaceous of Spain, exclusively of Central Asia: Zaraasuchus, Gobiosuchus, Zosuchus, and Artzosuchus from the Upper Cretaceous of Mongolia; Tagarosuchus from Lower Cretaceous of Southern Siberia; Edentosuchus, Sichuanosuchus, and Shantungosuchus from Lower Cretaceous of China. We report a new basal crocodyliform taxon, Neuquensuchus universitas gen.nov., sp.nov., from Neuquén Province, Argentina, belonging to Bajo de la Carpa Formation, representing the first and only “protosuchian” from the Cretaceous of Gondwana. The articulated and fragmentary materials belonged to a willowy, slender species, with very long and thin extremities. As in Shantungosuchus, the cervical centers are lengthened, with prominent ventral keel and well developed anteroventral parapophyses. As in basal crocodylomorphs, it possesses two sacral vertebrae. Also, a much enlarged scapular blade, with well developed acromial ridge and the posterior edge similar to Sichuanosuchus. The pronounced deltopectoral crest in the complete humerus is equivalent to Sichuanosuchus and as this, a circular, elongated and thin shaft with the medial condyle longer than the lateral one. Also, the complete ulna and radius is similar in their proportions to Sichuanosuchus. As this, the pubis is lengthened, very thin in the half section and not very expanded distally. The femur, tibia and fibula are elongated and similar to other non-derivated crocodyliforms. Besides representing the first Cretaceous “protosuchian” of Gondwana, the occurrence of these outside of Asia and Europe during the Cretaceous offers new evidence of pre-Albian dispersion between Gondwana and Central Asia through Europe. Key words: Crocodylomorpha. Protosuchian. Neuquensuchus universitas gen.nov., sp.nov. Cretaceous. Gondwana. RESUMEN: El primer “protosuquio” (Archosauria: Crocodyliformes) del Cretácico (Santoniano) de Gondwana. Los restos de “protosuquios” del Cretácico provienen, a excepción del “crocodyliforme de Las Hoyas” del Cretácico Inferior de España, exclusivamente de Asia Central: Zaraasuchus, Gobiosuchus, Zosuchus y Artzosuchus del Cretácico Superior de Mongolia; Tagarosuchus del Cretácico Inferior del sur de Siberia; Edentosuchus, Sichuanosuchus y Shantungosuchus del Cretácico Inferior de China. Aquí reportamos un nuevo taxón de crocodyliforme basal, Neuquensuchus universitas gen.nov., sp.nov., de la provincia de Neuquén, Argentina, correspondiente a la Formación Bajo de la Carpa, representando el primer y único “protosuquio” del Cretácico de Gondwana. Los materiales fragmentarios y articulados corresponden a una especie esbelta y delgada, con extremidades largas y delgadas. Al igual que en Shantungosuchus, los centros cervicales son alargados, con una quilla ventral prominente y parapófisis anteroventrales bien desarrolladas. Como en los crocodyliformes basales, Neuquensuchus posee dos vértebras sacras. Además, una hoja escapular muy expandida, con un puente acromial bien desarrollado y el borde posterior similar a Sichuanosuchus. La cresta deltopectoral pronunciada en el húmero es equivalente a la de Sichuanosuchus y al igual que este, la diáfisis es circular, alargada y delgada con el cóndilo medial mayor que el lateral. Asimismo, las proporciones del radio y la úlna son similares a Sichuanosuchus. Como este, el pubis es alargado, muy delgado en su sección media y poco expandido distalmente. El fémur, tibia y fíbula son alargados y similares a otros crocodyliformes no derivados. Además de representar el primer “protosuquio” cretácico de Gondwana, su presencia fuera de Asia y Europa durante el Cretácico ofrece nueva evidencia de un evento de dispersión pre-Albiano entre Gondwana y Asia Central a través de Europa. Palabras clave: Crocodylomorpha. Protosuquio. Neuquensuchus universitas gen.nov., sp.nov. Cretácico. Gondwana. 1 Submitted on September 14, 2006. Accepted on October 24, 2007. 2 Centro Regional de Investigaciones Científicas y Transferencia Tecnológica (CRILAR). Entre Ríos y Mendoza s/n, CP 5301, Anillaco, La Rioja, Argentina. E-mail: [email protected]. 3 Centro Paleontológico Lago Barreales (CePaLB), Universidad Nacional del Comahue. Ruta Provincial 51, km 65, Neuquén, Argentina. 418 L.E.FIORELLI & J.O.CALVO INTRODUCTION MATERIAL AND METHODS Fossil remains of basal non-Metasuchia The remains were found and gathered by Mr. Oscar Crocodyliformes from Cretaceous come almost de Ferrariis (at that time Director of the Museum exclusively from the Asian continent, to exception of the National University of Comahue), together of “Las Hoyas Crocodyliform” (SANZ et al., 1988) with J.O.C. The materials of this new basal (Fig.1) from the Lower Cretaceous of Las Hoyas, crocodyliform were originally referred as Spain (upper Barremian; DIÉGUEZ et al., 1995). Notosuchus (MUCPv-137) and were collected in The Asian forms are represented by species 1987. The study of the museum collection allowed coming from China, Mongolia and Russia. From us to find one more specimen represented by China comes Edentosuchus tienshanensis (YOUNG, fragmentary postcranial material but in good 1973; POL et al., 2004), a Protosuchia from the preservation (Fig.3). Lower Cretaceous of Tugulu Group, Xinjiang; Institutional abbreviations: GMPKU, Geological Shantungosuchus hangjinensis (WU et al., 1994) Museum, School of Earth and Space Sciences, Peking from the Luohandong Formation, Zhidan Group, University, Beijing, People’s Republic of China; IGM, Inner Mongolia and Sichuanosuchus shuhanensis Mongolian Institute of Geology, Ulaan Bataar, (WU et al., 1997) from an uncertain locality of Mongolia; IVPP, Institute of Vertebrate Paleontology Sichuan. From Mongolia come forms belonging and Paleoanthropology, Beijing, People’s Republic of to the Campanian age. Gobiosuchus kielanae China; LACM, Natural History Museum of Los Angeles (OSMÓLSKA, 1972; OSMÓLSKA et al., 1997) comes County, Los Angeles, California, USA; MACN, Museo from the Bayan Zak locality; Gobiosuchus Argentino de Ciencias Naturales, Buenos Aires, (?)parvus (EFIMOV, 1983), later considered Argentina; MUCP, Museo de Geología y Paleontología, conspecific of G. kielenae (OSMÓLSKA et al., 1997), Universidad Nacional del Comahue, Neuquén, comes from Üüden Sair locality; Zosuchus Argentina; UNC, Department of Geological Sciences, davidsoni (POL & NORELL, 2004a) and Zaraasuchus University of North Carolina at Chapel Hill; ZDM, shepardi (POL & NORELL, 2004b) come from Zos Zigong Dinosaur Museum, Zigong, Sichuan, China; Canyon locality; Artzosuchus brachicephalus ZPAL, Instytut Paleobiologii PAN, Warszawa, Poland. (EFIMOV, 1983), a very fragmentary form of uncertain filiation, comes from the same locality that G. (?)parvus. Lastly, Tagarosuchus kulemzini (ALIFANOV et al., 1999), with practically complete skull, comes from the Lower Cretaceous of Shestakovo locality, South Siberia. Here we present a new basal form of crocodyliform from the Upper Cretaceous of Northern Patagonia, Neuquén Province, Argentina. The remains come from the Bajo de la Carpa Formation, Neuquén Group (Fig.2), and represent the first “protosuchian” form for the Cretaceous of Gondwana. In this paper, we describe the anatomy of this new Crocodyliform together with a parsimony analysis of their Fig.1- Map of Eurasia showing the places of origin of the species of Cretaceous phylogenetic relationships. protosuchians. Arq. Mus. Nac., Rio de Janeiro, v.65, n.4, p.417-459, out./dez.2007 FIRST CRETACEOUS “PROTOSUCHIAN” FROM GONDWANA 419 Fig.2- Up right: satellital map showing the location of Argentina and Patagonia in South America; up left: satelital map of Northpatagonic region, showing the location of the Neuquén Province; below left: area of Comahue where were found and collected the materials of Neuquensuchus universitas, gen.nov., sp.nov. (scale bar = 10km - right inferior bar). Below right: stratigraphy of the Cretaceous of Neuquén Basin and stratigraphic column of the Neuquén Group (based on LEANZA et al., 2004. (Satellital images taken from GoogleEarth). RESULTS dinosaur Alvarezsaurus calvoi BONAPARTE, 1991 and Velocisaurus unicus BONAPARTE, 1991; GEOLOGY sauropod dinosaurs as cf. Laplatasaurus (LEANZA et al., 2004), Titanosauridae indet. (CHIAPPE & The Río Colorado subgroup constitutes the top of CALVO, 1994; pers.obs.), Neuquensaurus sp. the Neuquén Group; it is widely distributed in the (pers.obs.), Antarctosaurus and the peculiar South of the Neuquén Basin. The subgroup is beaked sauropod Bonitasaura salgadoi divided in two formations: Bajo de la Carpa (lower) APESTEGUÍA, 2004. Birds as Neuquenornis volans and Anacleto (upper) (LEANZA et al., 2004) (Fig.2). Bajo C HIAPPE & C ALVO, 1994 and Patagopteryx de la Carpa Formation is composed of coarse- deferrariisi ALVARENGA & BONAPARTE, 1992, snakes grained, light violet and pink sandstones of fluvial as Dinilysia patagonica WOODWARD, 1901, bird origin. The age has been dated as Santonian (LEANZA eggs in nests (SCHWEITZER et al., 2002), dinosaur et al., 2004) (Fig.2). eggs named Megaloolithus patagonicus CALVO et Outcrops in the area have given a wide variety al., 1997. Crocodyles are represented by of fauna such as carnotaurine abelisaurid Notosuchus terrestris
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